Invertebrate fossils from cave sediments: a new proxy for ... · sub-surface depositional processes...

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Biogeosciences

Invertebrate fossils from cave sediments: a new proxyfor pre-Quaternary paleoenvironments

O. T. Moldovan1, A. Mihevc2, L. Miko 3, S. Constantin4, I. N. Meleg1, A. Petculescu4, and P. Bosak2,5

1Department of Cluj, “Emil Racovita” Institute of Speleology, Clinicilor 5, 400006 Cluj-Napoca, Romania2Karst Research Institute, SRC SASA, Titov trg. 2, Postojna, Slovenia3European Commission, DG Environment, Av. de Beaulieu 5, 1160 Auderghem, Brussels, Belgium4“Emil Racovita” Institute of Speleology, Frumoasa 31, 010986 Bucuresti, Romania5Institute of Geology AS CR, v. v. i., Rozvojova 269, 165 00 Praha 6, Czech Republic

Received: 18 March 2011 – Published in Biogeosciences Discuss.: 30 March 2011Revised: 14 June 2011 – Accepted: 24 June 2011 – Published: 11 July 2011

Abstract. Five samples of clastic sediments from interiorcave facies taken in three Slovenian relic caves (Trhlovca,Raciska pecina, and a cave inCrnotice Quarry, Classi-cal Karst, SW Slovenia) provided invertebrate fossil re-mains. Most of them belong to Oribatida but sparse in-dividuals of Cladocera and insects were also identified.They represent the first pre-Quaternary invertebrate fossilsfound in sediments of continental temperate climate. ThePliocene/Pleistocene age of the sediments was determined bypaleomagnetic dating chronologically calibrated by micro-mammal biostratigraphy. Invertebrate fossils could be vali-dated as new proxy for the study of cave sediments due totheir suitability for ecological and paleogeographic correla-tions in caves and outside the caves. They also bring addi-tional information about cave formation and karst hydraulicregime in the area. Although the number of remains wasvery low, it is evidence that climatic conditions in caves al-low a better preservation of fossil remains of some groups ascompared to most of the surface habitats. This may open anew direction in the study of cave sediments.

1 Introduction

Cave sediments preserve the geological and paleoenviron-mental past (Horacek and Bosak, 1989) as well as bio-logical and anthropological information (e.g., Kukla andLozek, 1958; Horacek and Lozek, 1988; Bosak et al., 1989;

Correspondence to:O. T. Moldovan([email protected])

Sasowsky and Mylroie, 2004). This is of special impor-tance for the terrestrial (continental) history, where correl-ative sediments are mostly missing (Horacek and Bosak,1989), which is the case of the studied karst region. Cavesediments are formed in place in caves, or are allochthonousin origin (Kyrle, 1923; Kukla and Lozek, 1958). Two con-trasting facies can be distinguished among cave environ-ments (Kukla and Lozek, 1958). Theentrance faciesin-cludes fine-grained sediments transported from the vicinityof the cave by wind, water and slope processes. It repre-sents the most valuable section of the cave from a strati-graphic point of view as it may contain datable archeolog-ical and paleontological remains that are protected from sur-face erosion, weathering and biochemical alteration (cf. Fordand Williams, 1989, 2007). Theinterior faciesdevelops inthose parts of the cave that are more remote from the sur-face. A dominant part of sediments belonging to the interiorfacies form in vadose conditions. Sedimentary sequenceshere are extensive, consisting of fluvial gravels and sandsoverlain by flood or injecta deposits of laminated silts andclays often intercalated by speleothems. They may also con-tain dejecta, colluvial material and outer clastic sediments(including marine ones), often re-deposited and/or injectedfor longer distances within the cave (cf. Ford and Williams,1989, 2007). Most of the sediments result from material car-ried by (1) sinking streams and (2) gravity-driven infiltrationfrom the surface (e.g., Ford and Williams, 1989; Brinkmanand Reeder, 1995; White, 2007). Due to the dynamic en-vironment of cave interiors and periodicity of events, sed-imentary sequences often represent a series of depositionaland erosional events (sedimentary cycles). These are sepa-rated by unconformities (breaks in deposition), which may

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1826 O. T. Moldovan et al.: Invertebrate fossils from cave sediments

represent periods of substantial duration, and the erosionalphases may be of much longer duration that the depositionalevents (e.g., Bosak, 2002, 2008; Bosak et al., 2003). Suchsediments reflect past fluvial or lacustrine conditions withinthe cave. Distinct sequences of clastic sediments may becaused by changes in water volume and velocity, sedimentsource, surface weathering conditions and depositional con-ditions that are a function of climate change. Moreover, spe-cific topoclimatic features of the cave environments delay thedestruction processes of fossil remains (Sasowsky and Myl-roie, 2004; Polk et al., 2007). Therefore, many types of in-formation can be well preserved in the underground, suchas: (1) the above-ground source (which may no longer befound on the surface), (2) the environmental evolution on thesurface, including information on past vegetation and landuse which is not preserved in fast-changing surface soils(Kukla and Lozek, 1958; Bottrell, 1996; Courty and Val-lverdu, 2001; Panno et al., 2004; Polk et al., 2007), (3) thesub-surface depositional processes (Kukla and Lozek, 1958),and (4) the long-lasting general evolution of the sub-surfaceand surface karst (cf. Zupan Hajna et al., 2008b).

In karst areas, the evolution of the surface landscape isrecorded inside the caves; lowering of the base level shifts ac-tive speleogenesis to lower elevations, and the overlying pas-sages are subsequently abandoned and filled by sediments.Clastic sediments, sometimes intercalated with chemical pre-cipitates, transported from the surface through the caves arefrequently preserved unaltered for millions of years, provid-ing different types of information (Bosak et al., 1998, 2003;Sasowsky, 2007). Although the recognition that cave sed-iments can reflect and conserve paleoclimatic data is old(Kukla and Lozek, 1958 and literature herein), the scien-tific interest in paleoclimatic record from clastic cave de-posits developed only in the past decades (see summary ofWhite, 2007). Mostly fossil vertebrate remains have beenidentified in clastic cave sediments and used to character-ize environmental conditions, or used as paleoclimate prox-ies. Speleothems have been broadly utilized as paleoclimaticproxies using stable isotopes in combination with numeri-cal dating, palynology and sedimentological features of clas-tic cave sediments (Bastin, 1978; Bastin et al., 1986 amongothers). Environmental studies of cave sediments, especiallythose of the entrance facies, have been also carried out usingdifferent kinds of proxies; e.g., charcoal, sediment pollen andcoprolite pollen (Carrion et al., 1992a, b, 1995, 1997, 2005;Finlayson et al., 2008), stable carbon isotopes of organic mat-ter (Turney et al., 2001), pesticide content derived from agri-cultural pollution (Bottrell, 1996), the presence of a certainvegetation type (Panno et al., 2004), the fulvic acid fractionof the organic matter (Polk et al., 2007), magnetic mineralproperties (e.g., Elwood et al., 1996; Sroubek et al., 2001),or mollusks in cave archeological sites, etc. Other studiesthat focused on recent cave sediments as a proxy for the en-vironmental changes on the surface were specifically linkedto land-use (Polk et al., 2007). Only a single paper (Polyak et

al., 2001) mentioned the discovery of twelve species of ori-batid mites in two Holocene stalagmites from New Mexico,which were integrated in a paleoclimatic study.

Our study represents the first attempt to identify and studyfossil invertebrates in clastic sediments from the interior cavefacies and to discuss their possible use as biological proxiesin paleoenvironmental studies. Similar methods and proxiesapplied to other sedimentary deposits, such as lakes, seas andlotic environments, can also be applied to cave sediments inorder to obtain information on paleoclimate and paleoenvi-ronmental conditions at the time of sediment deposition. Sur-face sedimentary deposits, especially lacustrine ones, repre-sent paleoecological archives of plant macrofossils, pollen,algae and fossil invertebrates. Paleolimnology has devel-oped as a multidisciplinary science especially in the last twodecades using physical, chemical and biological proxies pre-served in lake sediments (Luoto, 2009). The structure and thecomposition of fossil assemblages (we prefer the term fossil,as proposed by Erickson and Platt, 2007, to “subfossil”) varyin response to changes in the environment, reflecting pastclimate, nutrient conditions, oxygen content, pH, pollutionor ecological interactions (Luoto, 2009). The cladocerans(Crustacea; Rautio, 2007), chironomids (Diptera; Walker,2001), and ostracods (Crustacea; Holmes, 2001) are the mostcommonly used invertebrate remains in paleolimnology. It isnot difficult to identify them to a species level, and their au-toecology is well known (Luoto, 2009). Other remains, suchas protozoans, bryozoans, oribatid mites (Acarina), insectsand mollusks, are rather rarely used (Smol, 2002). Until re-cently, invertebrate fossils have mostly been studied in lakesediments and only few of them in fluviatile environments(Gandouin et al., 2006; Engels et al., 2008; Howard et al.,2009).

Clastic sedimentary sequences belonging to the interiorcave facies were carefully selected for a first attempt to findfossil invertebrates in caves. Sedimentary sections well datedby magnetostratigraphy and paleomagnetic dates were cali-brated by biostratigraphy (Horacek et al., 2007; Zupan Hajnaet al., 2008b, 2010). The selected caves are located in theClassical Karst, the part of the Slovene Dinaric region ofKras (southeastern Europe). Shallow marine Dinaric Car-bonate Platform deposits (Jurassic to Paleogene) are coveredby Eocene flysch siliciclastics. Both units are overthrust intectonically complicated structures (Placer, 1999). No post-Eocene marine or terrestrial deposits are preserved on thesurface now. Surface morphology and karst evolved duringa single post-Eocene karst period. Speleogenesis was laterfollowed by cave infilling processes (partial or complete fos-silization) that started at the Oligocene/Miocene boundaryas indicated by fission track (AFTA) and paleontologically-calibrated paleomagnetic data from some of the studied sites(Zupan Hajna et al., 2010).

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O. T. Moldovan et al.: Invertebrate fossils from cave sediments 1827

Fig. 1. Location of the studied sites in Slovenia (see the geological map in the upper right corner; map after Plenicar et al., 1969): C = fossilcave in theCrnotice Quarry (the section is represented by an unroofed cave filled with yellow fluvial sediment covered by red clay withflowstone), T = Trhlovca Cave (the oldest part of the section with fluvial sediments), R = Raciska pecina Cave (the section is an alternationof flowstone and clay deposits; Photos by A. Mihevc).

2 Materials and methods

2.1 The studied sites

Three relic caves, all located in the Dinaric Classical Karst,were selected for this study – Trhlovca, Raciska pecina, anda cave inCrnotice Quarry (Fig. 1) – also due to the fact thatdetailed paleomagnetic dating was calibrated by micromam-mal biostratigraphy in two of them (Horacek et al., 2007).

Trhlovca Cave (45◦40′18.8′′ N; 13◦56′45′′ E) belongs tothe Divaska Cave System. The cave is part of an ancient andmore extensive system completely choked by sediments. Thecave was later partly rejuvenated and sediments exhumed as aconsequence of the evolution of the underlying Divaska Cave

(Bosak et al., 1998, 2000). The preserved fluvial sedimentsand speleothems, deposited in vadose conditions, are locatedin stratigraphically relevant position (Fig. 2b). A vertical sec-tion, 4.5 m in thickness, was described in detail by Zupan Ha-jna et al. (2008b). Its central part is well stratified and startsfrom the top with a brownish-red, clayey sand with intercala-tions of light-greyish and yellowish-brown sands. The rest ofthe section is represented by multi-colored clays in the upperhalf and chocolate-brown clays below. In the basal interval,silty to very fine-grained sandy admixtures occur in bandsand laminae. The arrangement of R (reverse) and N (nor-mal) polarized magnetozones shows ages older than 1.77 Ma(Zupan Hajna et al., 2008b).

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Table 1. A list of invertebrate fossils in the samples of cave sediments in Slovenia (only samples with identifiable animal remains are shown).

Site/Sample Short descriptionof the sediment

Identified taxa No.individuals

Observation

1 Trhlovca 1 (T1) red clay Miracarusn. sp. 1 new species

2 Trhlovca 2 (T2) beige clay Opiella (cf. Rhinoppia) n. sp. 1Dissorhinan. sp.

11

new speciesspecimen inpoor condition

3 Trhlovca 5 (T5) red clay Daphniasp.Dissorhinan. sp.Zygoribatula frisiaeTetramoriumsp.Insecta larvae

11111

new speciescosmopolitecosmopoliteunidentified

4 Raciska 4 (R4) red clay Oppiella(Rhinoppia) n. sp. 2Miracarusn. sp.Suctobelbellasp. ?

421

new speciesnew speciesspecimen inpoor condition

5 Crnotice 1 (C1) yellow clay OrthocladiinaeAstigmatida?

11

incompletespecimen inpoor condition

Raciska pecina Cave (45◦30’12.5′′ N; 14◦9’1.56′′ E) rep-resents a relic of an old cave system. The 2 m thick verti-cal section has a composite stratigraphic thickness of 6.5 m(Horacek et al., 2007; Zupan Hajna et al., 2008b; Fig. 2c).Its lower part consists of a vaulted stalagmite that includesseveral interbedded layers of red clays. It is overlain by athick interval of red clays with some silty and sandy interca-lations and thin calcite crusts and fossils (vertebrates and in-vertebrate –Potamon). The micromammals (withApodemus,cf. Borsodia) belong to middle to late Mammal Neogene bio-zone 17 (MN17; ca. 1.8–2.4 Ma; Horacek et al., 2007; posi-tion of R1 and R2 in Fig. 2). The clays are ponded and partlycovered by several collapse boulders. The upper part of thesection consists of subhorizontally laminated, porous andlight-colored flowstone with some ancient rimstone damsand interbedded red clays and silts. Lutitic interbeds betweenthe flowstone layers resulted from successive flooding thatdeposited well-sorted fine-grained allochthonous sediments.This may indicate either a distant position far from the ponorof the surface river or an allogenic stream passing througha system of sumps (Horacek et al., 2007). The top of thesection is flowstone with intercalations of brown cave loamswith bone fragments ofUrsus spelaeus. Fauna from the claypermitted the arrangement of the interpreted magnetozoneswith the Geomagnetic Polarity Time Scale (GPTS; Candeand Kent, 1995). The boundary of N- and R-polarized mag-netozone within the interval with fauna was identified withthe bottom of the C2n Olduvai subchron (1.770–1.950 Ma).The basal sediments can be correlated with the lower part ofthe Matuyama chron (2.150–2.581 Ma) and the Gauss chron(2.581–3.580 Ma) (Horacek et al., 2007).

Quarry operations in theCrnotice Quarry (45◦33’56.3′′ N;13◦52’47.7′′ E) uncovered many caves completely filled withsediments (Bosak et al., 1999, 2004; Mihevc, 2001, 2007;Fig. 2a). The studied section is located in the western quarrywall and represents a relic of an extensive, sediment-filledpassage with a diameter of about 10 m and a height of morethan 17 m. Its top is filled by speleothem breccia with redclay matrix. This is underlain by an interval, up to 4.5 thick,of light-colored, laminated silts and clays, sometimes sandy,overlying the rest of the section with deep erosion and a slightunconformity inside. The lowermost 7 m of the cave fill arecomposed of cyclically/rhythmically arranged multi-coloredfluvial sediments (clays to intraclastic microconglomerates).This fill rests on sessile tubes of the serpulidMarifugia cavat-ica on the northern cave wall (Mihevc, 2000; Mihevc et al.,2001). The mammal remains (withDeinsdorfiasp.,Bereme-dia fissidens, Apodemuscf. atavus, Rhagapodemuscf. fre-quens, Glirulussp.,Cseriasp.) that belong to MN15–MN16(ca. 3.0–4.1 Ma) were found in the same horizon as the ser-pulids (Horacek et al., 2007). The basal 1 m is composedof multi-colored laminated silts and clays developed in twosequences separated by an angular unconformity. The faunaindicated an age of the fill older than 1.77 Ma (base of theC2n Olduvai subchron). Most probably, the fill belongs tothe Gauss chron (2.6–3.6 Ma) or the other N-polarized sub-chron within the Gilbert chron (4.18–4.29 or 4.48–4.62 Ma;Bosak et al., 2004; Horacek et al., 2007).



Fig. 2. Lithological sections at the studied sites:(a) a sectionin the filled horizontal cave passage cut by theCrnotice Quarry(the original section): 1 = limestone, 2 = flowstone mixed with red-dish clay, 3 = large flowstone boulders, 4 = allogenic laminated flu-vial sediment, 5 = wall and sediment with tubes ofMarifugia ca-vatica, 6 = sampling points (C1–C2; modified from Bosak et al.,2004). (b) A section exposed in the Trhlovca Cave, several me-ters in thickness: 1 = limestone, 2 = reddish younger fill of clay,silt and fine sand, 3 = clays lighter in color, with a higher sandproportion, 3 = yellowish brown sandy clay, 4 = brownish to ochresandy clay, 5 = sampling points (T1–T5; modified from Bosak et al.,2006);(c)Raciska pecina Cave section: 1 = limestone 2 = flowstone,3 = red clay, silt and sand, 4 = brown clays with gravel and cave bearbones, 5 = stalagmites in flowstone, 6 = speleothems on the sectionsurface, 7 = collapsed limestone blocks, 8 = sampling points (R1–R4; modified from Zupan Hajna et al., 2008b).

2.2 Sampling protocols and invertebrate fossilidentification

A total of 11 samples were taken from the selected sites asindicated in Fig. 2. The samples were taken from the ex-posed faces, which had already been sampled for paleomag-netic and paleontological studies. Given the exploratory na-ture of our research, the samples were selected from what ap-peared to be distinct, stratigraphic units, located in positionsthat would allow a clear correlation with the previously es-tablished paleontological and magnetic chronostratigraphy.In the Crnotice Quarry, samples were taken only from theupper part of the originally described section that remainedintact after a recent collapse of the topmost sediments in thequarry face.

Fig. 3. Light microscope photographs of the identified fossil in-vertebrates in the cave sediments in Slovenia:(a) Daphnia sp.;(b) Orthocladiinae;(c) Dissorhinan. sp.;(d) Zygoribatula frisiate;(e) Miracarusn. sp.;(f) Opiella (cf. Rhinoppia) n. sp. 1;(g) Sucto-belbellasp. ?;(h) Oppiella(Rhinoppia) n. sp. 2.

Approximately 1 kg of sediment was taken from each sam-pling point and placed in sealed plastic bags with a label.In the laboratory, the samples were kept in 10 % KOH for30 min, and washed successively through sieves of 250 µm,125 µm and 40 µm. Sub-samples for each sieve dimensionwere examined separately under an Olympus SZX2 stere-omicroscope in 90◦ alcohol and each specimen was identi-fied under an Olympus BX51 microscope. Identification ofthe individuals was carried out following the specific meth-ods for each group.

3 Results and discussion

The number of identifiable invertebrate fossils in all sampleswas very low (Table 1). Some of the samples were com-pletely invertebrate-sterile. Unidentifiable animal fragmentsand vegetal fragments were also found. The only relativelywell-preserved specimens belong to the groups of Cladocera



(Crustacea), Oribatida (Acarina), and Chironomida and Hy-menoptera (Insecta). All these groups are commonly iden-tified in invertebrate fossil assemblages from lake sediments(Elias, 2007).

3.1 Faunal inventory

3.1.1 Crustacea Cladocera

OneDaphniasp. was identified in the Trhlovca Cave. Theidentified specimen is in a relatively good condition (Fig. 3a),but completely flattened laterally.Daphnia are known aslarge-bodied pelagic offshore cladocerans. Korhola (1999)and Korhola et al. (2000) found maximum lake depth to bethe most important factor explaining cladoceran distributionin Fennoscandian lakes. Jeppesen et al. (2001) found onlypost-abdominal claws, mandibles and ephippia ofDaphniaspecies in deep Quaternary lake sediments.

3.1.2 Acarina Oribatida

Mites of the suborder Oribatida (Acarina, Arachnida) aretypical soil-dwelling microarthropods, which can be alsofound in caves. Mites are of high potential value as bioindi-cators of the ecological conditions in terrestrial and aquaticecosystems (Lebrun and van Straalen, 1995; Behan-Pelletier,1999; Gulvik, 2007; Gergocs and Hufnagel, 2009). Almostall studies support the idea of a group with many represen-tatives living in humid habitats. Drought susceptibility ex-erts its effects via food limitation indirectly by a decrease ofmicrobiota in soil or other substratum as a result of lack ofwater. Nevertheless, some species adapted also to xeric con-ditions or high values of humidity. These minute arthropodsare usually preserved well enough in lacustrine or fluvial sed-iments and in sufficient numbers to be useful as proxies ininvestigations of Quaternary paleoclimate, paleoecology andstratigraphy (Solhøy and Solhøy, 2000; Solhøy, 2001; Polyaket al., 2001). Five species were identified in the studied sam-ples and four of them are new to science.

A new species belonging toMiracarus (Microzetidae)(Fig. 3e) was found in the sediments of the Trhlovca andRaciska caves. Modern representatives of the genus areforest-litter inhabitants.Oppiella (Rhinoppia)sp. 1 andOp-piella (Rhinoppia)sp. 2 (Fig. 3f, h), two species of the fam-ily Oppidae were identified in the sediments of the Trhlovcaand Raciska caves. Species of the genusOpiella (sensu lato)can be considered one of the most common arthropod groupson Earth (Norton and Palmer, 1991) with high diversity andabundance in forest litter, also present in shrublands, eco-tone zones and grasslands. The new species ofOppiellafrom the Raciska cave morphologically resembles a speciesknown from modern cave environments. The difference isin the length of sensilla, being much shorter than in the ex-tant form. The species is therefore an extinct element ofthe cave fauna of Slovenia.Dissorhinasp. (Fig. 3c) of the

same family (Opiidae) was represented in the Trhlovca Caveby two specimens, probably belonging to the same, newspecies. Some species of this genus prefer the border be-tween forest and open areas (Seniczak et al., 2006). Taylorand Wolters (2005) mentioned the tolerance of this genus todrought.Zygoribatula frisiae(Oribatulidae) (Fig. 3d) foundin the Trhlovca Cave is a species, found today in more aridsettings (Shepherd et al., 2002). The species is xero-tolerant,today known from repeatedly drying-out mosses and lichens,often in arboricol microhabitats.Suctobelbellasp. (Sucto-belbidae) (Fig. 3g) of the Trhlovca Cave resembles speciesof the recent genusSuctobelbellain some characters but dis-plays some specific characters, which may define a differentgenus. Species of this family are common in litter and upperorganic layer of forest soils, especially with a large amountof decomposing organic matter and with abundant fungal hy-phae. Some of the living species have been found in rottingwood and under bark of dead trees.

3.1.3 Insecta Chironomida

One representative of Orthocladiinae (Fig. 3b) was identi-fied in theCrnotice Quarry cave fill. The absence of men-tum made the identification of the lower taxon impossi-ble. Subfamily Orthocladiinae is a group of chironomidDiptera whose larvae prefer lotic habitats with cold and well-oxygenated waters (Dimitriadis and Cranston, 2001; Walker,2007). Representatives of this family inhabit cold and run-ning streams or unstable sandy bottoms of lakes, but are gen-erally adapted to low food (Walker, 2007) and are intoler-ant to low oxygen levels (Eggermont et al., 2008). This is awidely distributed family of chironomids and its representa-tives are frequently found in lake sediments.

3.1.4 Insecta Hymenoptera

One individual of the genusTetramoriumwas found in thesediments of the Trhlovca Cave. The genus is a typical inhab-itant of dry landscapes with shrubs. The individual lacks thehead, but is otherwise well preserved. It represents a genuswith very large distribution in the present fauna.

3.2 Paleoenvironmental significance of the fossilinvertebrates

Five taxa identified are new to science and their descrip-tion is in progress. Only a single taxon as yet identified ata species level is found in the modern fauna (Zygoribatulafrisiae). Two taxa were identified at a genus level (DaphniaandTetramorium) and one taxon as subfamily (Orthocladi-inae), due to their poor conservation. The cave richest infossils was Trhlovca Cave with six identified taxa in threeof the five samples. The maximum number of taxa belongsto the oldest sample (T5), presumably due to climatic condi-tions and sedimentation processes at the time of depositionand subsequently.



Fig. 4. A correlation of the obtained magnetostratigraphic resultswith the standard paleomagnetic scale (GPTS; after Cande andKent, 1995; modified after Horacek et al., 2007), mammal zones,temperature estimates (after Lisiecki and Raymo, 2005) and posi-tion of the samples with invertebrates (in red) and vertebrates (inblue): T = Trhlovca Cave, R = Raciska Cave, C = fossil cave in theCrnotice Quarry.

All the analyzed samples were taken from cave sedimentsdeposited during the Pliocene and Pleistocene periods, asdated by magnetostratigraphy and paleontological content(Fig. 4). During the Pliocene, the climate is globally knownto have become cooler and drier than during the Miocene(Robinson et al., 2008), followed by the more pronouncedPleistocene cooling. Broadly, two climatic stages are definedin the period covered by the deposition of clastic sediments

in the sampled caves (Fig. 5): the Early – Middle Pliocene(ca. 5.3 Ma to 3.6 Ma) with higher temperatures, and the LatePliocene – Early Pleistocene (ca. 3.6 to 1.77 Ma) with lowertemperatures, more similar to the present-day climate (Hay-wood, 2009).

Most of the invertebrate fossils were found in the TrhlovcaCave. Samples 1–3 belong to a more recent period than theolder infilling where samples 4–5 were collected, but all thestudied samples are older than 1.77 Ma (Zupan Hajna et al.,2008b, 2010). Clay samples T1 and T2 include taxa thatare typical for forest habitats with relatively high amounts ofdead organic matter in the upper horizon of soil (Miracarusand Oppiella) or taxa which can be found in the ecotonezones or in open, moderately humid areas (Dissorhina).From the oldest samples, T4–T5, only T5 contained fossil re-mains. The ecotone zone – typicalDissorhinawas found inassociation with dry habitat taxa, such asZygoribatulaandTetramorium. Fossils from these three samples indicate atransition from a warmer period with relatively drier vegeta-tion (Early or Middle Pliocene) to a more humid and forestedhabitats of the Late Pliocene/Early Pleistocene. The presenceof Daphniasp. in sample T5 suggests the presence of a lakeor a low hydraulic regime of the stream, as well as the prox-imity of cave entrance. This assumption is also supported bythe varved sediments that indicate continuous deposition inthe past, during single-flood events or flood pulses that prob-ably lasted less than a few thousand years (Zupan Hajna etal., 2008a). The relatively good preservation of this clado-ceran is also indicative of a slow flow from the surface downto place of deposition, possibly due to the still epiphreaticposition of some parts of the subterranean system (Mihevc,2007) and to the short-distance transport. The presence ofDaphniaspecies, as an indicator of oligotrophic (low food)environments (Szeroczynska and Zawisza, 2005), supportsthe hypothesis of a drier vegetation on the surface.

Only sample R4 provided fossil invertebrates in Raciskapecina. These are represented by three individuals of orib-atid mites belonging to the same ecological group of forestinhabitants. Sample R4 comes from the upper half of thesection of this cave that belongs to the Early Pleistocene (ter-mination of the Olduvai-chron).

The only chironomid identified at a family level was foundin sample C1 inCrnotice Quarry. Located in the upper halfof the sediment section, the Orthocladiinae dipteran can beused as an indicator for colder and well-oxygenated waters.The individual was carried underground by a relatively fastflowing river, which suggests a colder climate with more in-tense precipitation towards the end of the Pliocene.

3.3 Correlations with other proxies

The discovery of five new species (and perhaps a new genus)is important not only for the taxonomy of the group ofmites (in this case), but is also of paleoenvironmental sig-nificance. The Pliocene/Pleistocene cave sediments show



Fig. 5. Invertebrate fossils found in the studied caves, correlated with climate, vegetation and fossil vertebrates.

a very low concentration of taxa, or none at all in morethan half of the analyzed samples. The scarcity of faunaremains makes further biostratigraphic interpretation impos-sible, but the presence and ecology of some taxa can becorrelated with the discovered vertebrate remains as well aswith the climate and corresponding vegetation of the differ-ent Pliocene/Pleistocene periods, indicating a transition froma dry or mild climate with mixed forest to a colder climateand more humid forest (Table 2, Fig. 5). The use of less fre-quently occurring zoological remains in paleolimnology wasemphasized by Luoto (2009) as an important proxy for envi-ronmental changes.

The fossil vertebrate assemblage from theCrnoticeQuarry, includingDeinsdorfia sp., Beremendia fissidens,Apodemuscf. atavus, Rhagapodemuscf. frequens, Gliru-

lus sp. andCseriasp. (Horacek et al., 2007), is typical fora temperate and humid climate. The high number of for-est and shrub inhabitants (Apodemus, Glirulus, Myodesgen-era), together with taxa that are widely distributed in temper-ate and humid climates (Beremendia fissidens, BlarinoidesandDeinsdorfiagenera), is indicative of a mild climate. AtRaciska pecina Cave, the most important difference is thepresence ofMyodesandBorsodia(Horacek et al., 2007) asindicators of a colder and drier climate. Owing to the scarcepresence of identifiable invertebrates, a correlation with theequally scarce vertebrate record is problematic. Only threeof the invertebrate samples (C1, T1 and R4) come fromthe same periods as the found vertebrate fossils, and cor-relations were made only for these samples. Sample C1provides a single indicator for cold/well-oxygenated water



Table 2. Taxa found in cave sediments of Slovenia with the corresponding vegetation, sediment type and origin, and vertebrate fossils.

Stage TAXA Environment Sediment Vertebrates

Trhlovca Raciska Crnotice origin

UpperPliocene/LowerPleistocene

MiracarusOpiella Dissorhina

– – forest,ecotone zones

surface,fluvial

–

– Oppiella MiracarusSuctobelbella(Potamon∗)

– forest,ecotone zones,grasslands + river

surface,fluvial

(fish teeth∗),micromammalsliving in steppeand tree habitats

– – Orthocladiinae(Marifugia cavatica∗)

cold,well oxygenatedwater

surface,fluvial

(fish teeth∗),micromammalsliving in treeand riversidehabitats

LowerPliocene

Daphnia,Dissorhina,ZygoribatulaTetramolium

– – dry landscape withshrubs + river

surface,fluvial

–

∗ Mentioned in Horacek et al. (2007).

(Orthocladiinae), which is supported by the presence ofBere-mendia, Rhagapodemustypical for temperate climate, and ofCseriaandMimomysthat are water-related micro-mammals.Samples T1 and R4 include Oribatida invertebrates (generaOppiella, Miracarus and Suctobelbella) that inhabit forestlitter, shrublands, ecotone zones and grasslands, and verte-brates which indicate the presence of trees in wet forests andsteppic habitats, typical habitats forMyodesandBorsodia.The presence of invertebrate drought-tolerant taxa in T5 in-dicates a landscape with shrubs and grassland during the lateEarly Pliocene. The presence of forest inhabitants in T1 andR4 correlated with the presence of cold/dry-associated verte-brates points to a colder climate during the Mid-Pleistocene.Between these two chronologically extreme samples, inver-tebrates were also identified in samples C1 and T2. Sam-ple C1 corresponds to the advent of the Late Pliocene cool-ing and probably higher flow rates on the surface, which ex-plains the presence of the chironomid. Sample T2 fits withinthe picture of forests mixed with open areas during the EarlyPleistocene.

No pollen was found in the studied samples and sec-tions, but detailed studies on the nearby Italian and peri-Mediterranean sites given in Bertini (2010) can be well cor-related with the invertebrate fossils found in the Sloveniancave sediments (Table 2). During the Zanclean, the cli-mate was warmer and drier or more humid than today, de-pending on the geographical latitude and altitude (Faque-tte et al., 2006); forest taxa of humid subtropical to warmtemperate zones were dominant (Bertini, 2010). The gen-

eral decrease of humidity and installation of drier conditionsduring the Piacenzian occurred prior to 2.5 Ma (Bertoldi etal., 1989). Alternations of forest vegetation and steppes,with typical open vegetation phases characterized the fol-lowing early Pleistocene, during the Gelasian and the Cal-abrian (Bertini, 2010). In general, the floral compositionand structure of vegetation of the Pliocene-Pleistocene arecharacterized by the rare appearance of new species and byspatially asynchronous events of disappearance of the sametaxon (Bertini, 2010). Frequent changes in both climate andvegetation can explain the extinction of some of the identi-fied invertebrate taxa in the cave sediments of Slovenia. Ingeneral, faunal changes depend on the environment, definedmostly by climate and vegetal associations. If changes arefrequent and of relatively short duration, some of the taxa thatare unable to adapt may get extinct, or evolve into differentspecies. Oribatida is a group with a lower mobility than fly-ing invertebrate species, and their migration can only be overshort distances, following the slowly migrating vegetation orvegetal associations. The identified new Oribatida speciescan be considered either members of an extinct faunal groupwith no relatives in the present fauna of southern Slovenia, orancestors of a living species which have undergone evolutionin the present-day habitats or in different areas nearby. Veg-etation succession in time contributed to the settlement of acomplex mosaic of different climates/biomes in the Mediter-ranean zone (Bertini, 2010). This situation documented forflora is supported by the invertebrates found in the cave sed-iments. The section that provided most of the specimens,



in the Trhlovca Cave, yielded taxa indicative of mixtures offorests with open dry areas. As already documented by Fin-layson et al. (2008) for more recent sediments, environmen-tal changes must be taken into account at small or mediumscales. Invertebrate fossils can provide information at smallscales and must be associated with other proxies and prefer-ably with more sites of the same region in order to providepaleoenvironmental information that would be significant atregional scale.

3.4 Implications for the reconstruction of karstevolution

If the standing condition of the fossil is known, its endpointcan be used to infer some of the geological processes thatacted on the body during its transport to the site of deposi-tion (Erickson, 1988). The presence of surface animals inpre-Quaternary interior facies of cave sediments is indicativefor an intense karst evolution of the Dinaric karst, includ-ing the filling of a part of the cavities with sediment duringthe Neogene. The very low abundance of fossils depositedin clastic cave sediments of the interior facies is well known(e.g., Horacek and Lozek, 1988). This is probably relatedto the runoff in the drainage area or to the hydraulics ofthe subterranean streams. Almost all of the studied inver-tebrate samples have at least one aquatic component, and all(with a single possible exception) are surface invertebrates,which were brought inside the caves by allogenic streamsand deposited together with the sediments. The most com-mon clastic sediments in the studied caves are different typesof fine laminated clays and silts. They were deposited fromfloodwater suspended load under conditions of pulsed flowor cave lakes comparable with slackwater facies of Boschand White (2004). This depositional process corresponds tomore or less regular flooding of karst areas by sinking rivers.Mineral assemblages of the cave deposits were derived fromhighly homogenized weathering products of Eocene flyschsediments and soil cover on Paleocene and Cretaceous lime-stones (Zupan Hajna et al., 2008b). The tectonically-drivenlowering of the regional base level was connected with thechange in tectonic regime at ca. 6 Ma (Vrabec and Fodor,2006). This caused a transition from an epiphreatic to a va-dose regime, followed by a decrease in hydraulic head insome cave systems or their higher-situated parts and a com-plete fill with fluvial sediments as a consequence (Mihevc,2007; Zupan Hajna et al., 2008b).

As already mentioned, the presence ofDaphnia sp. andits relatively good preservation in sample T5 indicates theexistence of a system with low hydraulic head at the cave en-trance and multiple flood events that contributed to the con-tinuous deposition of varved and/or cyclically arranged sed-iments. The chironomid in sample C1 indicates a period ofrelatively high flow rate in a vadose regime. The chironomidwas found at the same level asMarifugia cavatica, serpulidpolychaetes attached to walls at the air/karst-water interface,

still living in the caves of the Dinaric karst (for summary seeMihevc et al., 2001).

Pre-Quaternary arthropod remains, abundant in marinesettings and quite common in amber, are rare and have yetbeen reported in continental sediments only from Arctic en-vironments (see a complete list in Elias, 2010). The deposi-tional mechanisms and the low intensity of biochemical pro-cesses can explain the relatively good state of preservationof old invertebrate remains both in the cave sediments andArctic lake sediments. Although the number of identified in-vertebrates at the studied Slovenian sites is small, their stateof preservation is relatively good considering the age of thesediments. This may suggests a combination of: (1) a rela-tively short and slow transport to the site of deposition, (2) arapid burial, i.e., a high sedimentation rate, and (3) subduedmicrobial and biochemical processes that could have alteredthe entire organisms.

4 Conclusions

Three sites of relic caves from the Classical Karst of Sloveniaprovide first evidence of Pliocene/Pleistocene invertebrateremains in continental clastic sediments from interior cavefacies in temperate regions. These finds suggest that cavesediments can preserve yet another proxy for the assessmentof paleoclimatic and paleoenvironmental conditions. Thisnew proxy may be important for a number of reasons: (i) thescarcity of vertebrate fossils in pre-Quaternary cave deposits;(ii) the need for cross-validation of inferred paleoenviron-mental settings in the case of incomplete or ambiguous co-eval proxies; (iii) the cave conditions that are prone to a betterpreservation of old chitinous invertebrates when comparedto surface settings; (iv) its added-value as a source of in-formation on speleogenesis and evolution of karst hydraulicregimes even for paleokarst settings.

Caves are known as systems with low-energy input fromthe surface and low-energy in situ production. This mayhamper the use of fossil invertebrates from caves due tothe relatively low chance of finding identifiable cave orsurface specimens transported into caves. However, forthe three described sites, a relatively low number of in-vertebrate specimens were successfully identified and datedby biostratigraphy-calibrated magnetostratigraphy back toca. 1.77 to ca. 4.8 Ma, i.e., across the Pliocene/Pleistoceneboundary. When cross-correlated with the coeval vertebratefauna, these findings support the idea of a colder phase as-sociated with the Pliocene/Pleistocene transition, while alsobringing new insights on the regional karst evolution.

Acknowledgements.The authors are grateful to Ilse Bartsch,Karina Battes, Steve Brook, Mirela Cımpean, Balint Marko,Laura Momeu for the help with taxa recognition and ecologicalinformation. Jirı Adamovic has kindly helped with the Englishrevision of the manuscript. This study was funded through the



KARSTHIVES Project PCCEID 31/2010 funded by CNCSIS-UEFISCDI (Romania; granted to SC), Grant Agency AS CRNo. IAA300130701 (Paleomagnetic research of karst sediments:paleotectonic and geomorphological implications; granted to PB)and the Institutional Research Plan No. AV0Z30130516 of the GLIAS CR, v. v. i. The authors thank to editor Wolfgang Kiesslingand reviewers Ira Sasowski and John Holsinger for their insightfulcomments.

Edited by: W. Kiessling

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